Introduction You have studied the salient features and classification of Protochordata and the salient features of the

subphyla Hemichordata, Urochordata and Cephalochordata included under the group Protochordata in units 1 and 2. Aspects of morphology, anatomy, physiology and affinities of Balanoglossus, a type representing Protochordata are explained in unit 2. You will learn the salient features, classification and an account of typical examples representing the main groups of the subphylum Vertebrata from unit three onwards. Unit three presents the salient features, and classification of the earliest group of Vertebrates, The Agnatha. A detailed description of Petromyzon, as an example of this group is given. Agnatha are distinct from other Vertebrates in the absence of jaws and paired fins or appendages. Hence, Agnatha are considered to be the most primitive vertebrates. Agnatha Salient features The term Agnatha refers to the vertebrates without jaws (a not; gnathos jaw). Agnatha includes primitive, fish like vertebrates, having an elongated body. Notochord persists throughout life. Incompletely developed vertebrae are present over the notochord. Dorsal nerve cord is present. Its anterior region is differentiated into a brain. Pharyngeal gill slits range from 5 to 16 pairs. Mouth is circular, ventral in position and suctorial type. Paired fins are absent, while median and tail fins are present. A distinct head region is present, with a single, median nostril and with a pair of eyes. Auditory organ contains only two semicircular canals. Skin is soft and smooth, containing mucous glands. Scales are absent. Endoskeleton is cartilaginous in nature. No bony structures. Heart two chambered, with one auricle and one ventricle. Blood with nucleated erythrocytes and leucocytes. Sexes are separate. A single gonad is present without a gonoduct. Marine or freshwater forms. Fertilization is external. Development direct or indirect which includes a larval stage. Self Assessment Questions I Choose the correct answer: 1. In Agnatha, the heart is a) Non-contractile b) Single chambered c) Two chambered d) Three chambered 2. Endoskeleton of Agnatha is made of a) Bone b) Cartilage c) Bone and cartilage d) Membrane State whether true or false: 3. In Agnatha, anterior region of the nerve cord is differentiated into a brain. true 4. The auditory organ contains three semi-circular canals.false

Agnatha Classification The division Agnatha is represented by about 50 species of the living jawless vertebrates. A group of extinct primitive jawless vertebrates called Ostracoderms are also included under this division. Agnatha is divided into two classes namely Ostracodermi and Cyclostomata. Class 1. Ostracodermi: The name Ostracodermi refers to the shelled skin or bony skin of these jawless vertebrates (Ostrakon shell; derma skin). The bony plates or scales form a heavy bony armour as a protective exoskeleton. Fish like vertebrates without jaws. Ostracoderms are known to be the most primitive fossil vertebrates. All members have become extinct and the group is represented by fossils only. No paired fins but with a pair of spines, behind the gill region. Mouth slit-like. A single nostril on top of the head. A single pineal eye present. Head and gill regions are covered with a bony shield. Some members possess internal bony-skeletal structures. Eg: Cephalaspis Hemicyclaspis

Fig. 3.1: Hemicyclaspis Class 2. Cyclostomata: The name Cyclostomata refers to the circular mouth of these jawless vertebrates (cyklos = circular; stoma = mouth). Body fish-like, elongated and eel like. Notochord remains throughout life. Anterior region of nerve cord is differentiated into a brain. Pharyngeal gill slits ranging from 5 to 16 pairs. Mouth is circular, ventral or terminal in position and suctorial. Paired fins are absent. Median and Caudal fins are present. A distinct head region with a median nostril and a pair of eyes. Skin is soft, smooth and contains mucous glands. Endoskeleton is cartilaginous. Cyclostomata is called by another name Marsipobranchi, due to the presence of pouched gills (gills in sac like pouches). Marine or fresh water animals, having a wide distribution. Probably, the degenerate survivors of the fossil Ostracoderms. This group is represented by the lampreys, hagfishes and slime eels.

Fig. 3.2: Myxine (Hagfish) Class cyclostomata is divided into two orders namely Petromyzontia and Myxinoidea:

circular mouth is present at the center of the buccal funnel. It encloses a protrusible tongue, bearing large horny teeth. This rasping tongue serves to remove and feed on the tissues of the host fish. Thus, the buccal funnel serves for attachment to the host and in feeding. The head bears a pair of lateral eyes and a single nostril. The eyes lack eyelids. The nostril also called naso-hypophyscal aperture is mid dorsal in position. On each lateral side of the head, there are seven gill-slits, behind the eye. The anus lies at the base of the tail, on the ventral side and behind the anus is the cloacal aperture or urinogenitial aperture. Numerous sensory pores are present on each lateral side of the body, representing the lateral line sense organs.

Self Assessment Questions II State whether true or false: 1. The order Petromyzontia includes animals commonly called the hag fishes. false 2. The class Ostracodermi includes only extinct forms.true Fill in the blanks: 3. The name Cyclostomata refers to the presence of circular mouth_. 4. The fossil Agnatha are collectively called Ostracoderms 3.4 Petromyzon Type study Petromyzon is a jawless vertebrate and hence is included in the division Agnatha. Due to the presence of a circular mouth, it is included under the class Cyclostomata. Petromyzon is commonly called the lamprey. 3.4.1 Habitat and Habit Petromyzon marinus commonly called the sea lamprey is a marine form, having wide distribution. The fish like adult Petromyzon lives in the sea and migrates to rivers for spawning in the spring seasons. It is parasitic on fishes, turtles and other marine animals. It has a circular and suctorial mouth surrounded by numerous horny teeth in a special region called buccal funnel. The horny teeth and rasping tongue help in clinging to the host and for feeding respectively. The development includes a larval form which is a fresh water, filter feeding organism.

Fig. 3.4: Buccal funnel region of Petromyzon 3.4.4 Body wall and muscles The skin is soft and smooth without scales. It consists of an upper epidermis and a lower dermis. The epidermis consists of mucous glands secreting a slimy mucous. The dermis consists of collagen and elastin fibres and a sub-cutaneous tissue with blood vessels, fat tissue and connective tissue. E-shaped segmental muscles called myotomes are present in the trunk and tail regions. This powerful musculature helps in locomotion. The sucking action of the buccal funnel and the movement of the rasping tongue are controlled by special muscles. 3.4.5 Endoskeleton In Petromyzon, the endoskeleton is cartilaginous. True bone is absent. The notochord and skull form the axial skeleton of the animal. Notochord remains throughout life. The skull consists of cranium which encloses the brain and sense capsules which enclose the sense organs. Presence of a branchial basket or visceral skeleton is a special feature. This structure consists of nine vertical bars and four pairs of transverse bars made of cartilage. This is not homologous with the visceral arches of fishes. This branchial basket supports and protects the gill pouches. A thick bar called lingual cartilage is present in the tongue, giving support to it.

Fig. 3.3: Petromyzon marinus 3.4.2 External morphology The adult has an elongated body which is divisible into head, trunk and tail regions. The body is cylindrical, while the tail is laterally compressed. The colour of the animal is greenish-brown. The body surface is smooth without scales and contains mucous glands for production of a slimy secretion. Petromyzon marinus attains a length of about one metre. Paired fins are absent. It has one or two dorsal fins and a caudal fin. 3.4.3 Buccal Funnel The head bears a cuplike structure called buccal funnel or sucker. This sucker contains an outer membrane called marginal membrane, with many small projections, the oral fimbriae. These projections serve for attachment to the host. Many sensory structures called cirri are present, projecting from the marginal membrane. The buccal funnel consists of conical, yellowish horny teeth arranged in rows. A small,

Fig. 3.5: Cartilaginous skeleton of Petromyzon 3.4.6 Digestive System The mouth is present in the buccal funnel. Jaws are absent. The buccal funnel or sucker is used for attachment to the body of a fish or other host animals. The mouth leads into a buccal cavity. The rasping tongue with horny teeth is protrusible and by its movement, can open or close the mouth. The buccal cavity leads into two tubular structures, a dorsal oesophagus and a ventral respiratory pharynx. This is the pharyngeal region which ends in a respiratory tube. This tube contains seven internal gill slits on each side. Presence of this respiratory tube is a special feature of Petromyzon. The narrow tubular oesophagus leads into a straight tube, the intestine. The latter contains a spiral valve, having a role in digestion and absorption. The intestine leads into rectum which opens outside by anus in the cloacal region. The digestive

glands include a pair of salivary glands or buccal glands and a large bilobed liver. These buccal glands secrete an anticoagulant substance. A gall bladder and a bile duct are present in the larva but are absent in the adult. Petromyzon is adapted for a parasitic mode of feeding. The buccal funnel serves for attachment to the host fish. The rasping tongue serves to remove small pieces of tissues of the host animal. The anticoagulant secretion injected into the wound prevents clotting of blood. Petromyzon sucks the blood and body fluids of the host animal. 3.4.7 Respiratory system There are seven gills, located in pouches called gill pouches on each side. These pouches lie in between the body wall and the respiratory pharynx. Each gill pouch contains a series of gill filaments or gill lamellae, supplied with blood capillaries. The gill pouches open outside by external gill slits. Normally water is drawn through the mouth into the gill pouches both in the larva and adult. Exchange of gases takes place in the gills and water escapes through the external gill slits. When the adult is attached to a host by buccal funnel, water enters the gill pouches and leaves the gill pouches only through the external gill slits. A valve like structure called velum present at the entrance of the respiratory pharynx prevents the return of water from the pharynx to the buccal cavity. 3.4.8 Circulatory System This system consists of heart, arteries and veins to supply and collect blood. The heart is simple, ventrally located behind the last pair of gill pouches and is covered by a pericardium. It is two chambered, with an auricle and one ventricle. It has sinus venosus. The venous blood alone circulates through the heart; the circulation is called single circulation. This type of heart is called venous heart. Ventricle pumps the venous blood into a ventral aorta. This vessel gives off seven pairs of afferent branchial arteries taking the blood to the gills. The oxygenated blood is collected through seven pairs of efferent branchial arteries which open into a median dorsal aorta which supplies the oxygenated blood through numerous arteries to the tissues. The venous blood is collected from the anterior and posterior regions of the body through the veins called anterior and posterior cardinal veins. Blood contains erythrocytes with hemoglobin and leucocytes. 3.4.9 Nervous System This system includes a simple brain, cranial nerves and spinal nerves. The brain is of primitive type, having a pair of cerebral hemispheres, a pair of optic lobes, a reduced cerebellum and medulla oblongata which is continued into a spinal cord. The diencephalon bears the pineal apparatus dorsally and the pituitary body and infundibulum ventrally. The brain encloses cavities called ventricles. Ten pairs of cranial nerves are present. Spinal nerves with separate dorsal and ventral roots are present. 3.4.10 Sense Organs A pair of lateral eyes is present without eyelids. A single mid dorsal nasal aperture leading into a small olfactory sac is present in the head. The auditory organ consists of two semicircular canals. Lateral line sense organs are present on the lateral sides of the body. 3.4.11 Excretory System Pronephros type of kidneys are present in the larva of Petromyzon. The adult has a pair of mesonephric type of kidneys. The nephron or uriniferous tubule contains glomeruli inside the bowmans capsule. The nitrogenous wastes from the blood are collected in glomeruli and are eliminated through the ureter into the urinogenital sinus.

Fig. 3.6: Petromyzon (A) External features; (B) Longitudinal section 3.4.12 Reproduction and development Sexes are separate. A single large gonad is present, gonoduct is absent. Mature spermatozoa from males and eggs from females are released into the coelom and then into the water through urinogenital sinus. Mature adults migrate to rivers for spawning in spring. The male builds a nest as a shallow depression. Spermatozoa and ova are released into the nest. Fertilization occurs in the nest. The fertilized eggs hatch out in about three weeks into a larva called ammocoete larva. It is a minute, transparent and free swimming larva showing resemblance to Amphioxus. The larva has a single, continuous median dorsal fin. It is a filter feeder and non-parasitic. Sucker and teeth are absent. Presence of a tubular endostyle, peripharyngeal grooves, epipharyngeal grooves, velum and many gill slits are the important features of the larva and in having these features, the larva resembles Amphioxus. Feeding and respiration are similar to those of Amphioxus. The larval period lasts for a few years, ranging from 3 to 7 years. The free swimming, filter feeder undergoes metamorphosis into a semi parasitic adult lamprey. Several structural changes occur during metamorphosis. The young lamprey migrates into the sea, where they attain sexual maturity.

Fig. 3.7: Ammocoete larva of Petromyzon 3.4.13 Affinities Petromyzon and other cyclostomes represent the most primitive jawless vertebrates. They possess the basic and distinguishing features of chordates. Cyclostomes are considered to be the last degenerate survivors of their closely related fossil Ostracoderms. The Ammocoete larva of Petromyzon resembles Amphioxus in having several similar characteristic features. However, cyclostomes show many advanced features such as the presence of a distinct head with sense organs, welldeveloped brain and nerves, well developed circulatory system, cartilaginous endoskeleton and body wall. In having these features, Petromyzon and other cyclostomes resemble fishes and other higher vertebrates. Presence of a buccal funnel with horny teeth, rasping tongue, respiratory pharynx with gill pouches and branchial basket are the specialized features of Petromyzon.

Self Assessment Questions III Choose the correct answer: 1. The return of water from the respiratory pharynx to buccal cavity is prevented by a valve called velum. a) Spiral valve b) Velum c) Atrial valve d) Buccal valve 2. In Petromyzon, the respiratory structures are located in the gill pouches. a) Pharynx b) Buccal cavity c) Gill pouches d) Branchial basket State whether true or false: 3. Ammocoete larva is a filter feeding organism.true 4. Endoskeleton of Petromyzon is made of bone.false 5. In Petromyzon, paired fins are absent.true 3.5 Summary This unit explains the salient features, and classification of Agnatha using Petromyzon as an example. The study reveals that Agnatha are the earliest and most primitive vertebrates without jaws and paired fins. Agnatha is divided into two classes namely Ostracodermi and Cyclostomata. Petromyzon and other cyclostomes represent the most primitive jawless vertebrates. They possess the basic and distinguishing features of chordates. Presence of a buccal funnel with horny teeth, rasping tongue, respiratory pharynx with gill pouches and branchial basket are the specialized features of Petromyzon.